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This Work Is Licensed Under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License. to View a Copy This work is licensed under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-sa/3.0/us/ or send a letter to Creative Commons, 171 Second Street, Suite 300, San Francisco, California, 94105, USA. REVISION AND CLADISTICS OF THE MIDDLE AMERICAN GENUS CREAGROPHORUS MATTHEWS (COLEOPTERA: LEIODIDAE) QUENTIND. WHEELER Department of Entomology The Ohio State University Columbus, OHIO 43210 Quaestiones Entomologicae U.S.A. 15:447-479 1979 Creagrophorus is a compact genus of four Middle American species whose members eat fruiting bodies of puffballs (Gasteromycetes); Creagrophorus hamatus Matthews (Mexico, Panama), Creagrophorus bihamatus Matthews (Panama), Creagrophorous jamaicensis Peck (Jamaica), and Creagrophorus spinaculeus new species (Panama). A cladistic analysis led to the following conclusions: Creagrophorus is supported as a monophyletic group by apotypic broad labial palpus segment HI with taeniaform seta, spines and flagellum of endophallus, absence of epicranial lines, and fimbriae of galeae and presence of four setae on segment I of the urogomphus of the larvae, transverse coxites and flattened, sclerotized styli of female genitalia, male femoral hooks (denticles), expanded middle tibia, and puffball ecological relationships; the Aglyptinus association is a monophyletic lineage consisting of Creagrophorus, Aglyptinus, and Scotocryptini, and defined synapotypically by 3-3-3 tarsi and oblique lines on abdominal sternum HI; Aglyptinus is the sister group of Cre&grophorus-Scotocryptini; Scotocryptini is the sister group of Creagrophorus; C. hamatus is the sister to the jamaicensis-bihamatus-spinaculeus lineage, and C. bihamatus sister species to C. spinaculeus. Creagrophorus is hypothesized to be an endemic Middle American genus, which arose after its ancestor migrated from South America where the sister group differentiated in body structure and ecological relationships (Scotocryptini); the hamatus lineage was isolated north of the Isthmus of Tehuantepec; the jamaicensis lineage was isolated in nuclear Middle America (and dispersed to Jamaica); and bihamatus/spinaculeus became north/south vicariants due to volcanism in the Chiriqui region. Creagrophorus beetles are continental in differentiation pattern, and as such are implied to have wide geographic ranges and require relatively long periods of time for speciation. The Aglyptinus association and Creagrophorus are defined, species of Creagrophorus described, and the following presented; keys to genera o/Aglyptinus association and species of Creagrophorus, habitus drawings of adult and larval beetles, illustrations of important character states and geographic distribution, designation of lectotypes for Matthews' species, and discussions about ecological, cladistic, and zoogeographic relationships. Creagrophorus est un petit genre de Leiodidae d'Amerique Centrale comprenant quatre especes qui se nourrissent de vesses-de-loup (champignons Gasteromycetes): ce sont: C. hamatus Matthews (Mexique et Panama), C. bihamatus Matthews (Panama. C. jamaicensis Peck (Jamaique) et C. spinaculeus Wheeler, une espece nouvelle (Panama). Une analyse cladistique permet de tirer les conclusions suivantes; les Creagrophorus forment un taxon monophyleetique, a cause de la presence des caracteres apotypiques suivants: troisieme segment des palpes labiaux large et portant une soie taeniaforme, phallobase avec epines et flagelle, absence de lignes epicraniales et de fimbriae sur les galeas, presence de quatre soies sur le premier segment de I'urogomphe chez les larves, coxas transversaux et aplatis, stylets de I'armature genitale des femelles sclerifies, femurs des males portant des crochets (denticules), tibias intermediates elargis, et rapports ecologiques avec les vesses-de-loup: le groupe Aglytinus constitue une lignee monophyletique et comprend les Creagrophorus, les Aglyptinus et les Scotocryptini. Ce groupe est defini par les synapomorphies suivantes: formule tarsale 3-3-3. et presence de lignes obliques sur le troisieme sternite abdominal. Le genre Aglyptinus est monophyletique et est apparente aux CTeagrophorus-Scotocryptini. C. hamatus est apparente a la lignee jamaicensis-bihamatus-spinaculeus et C. jamaicensis est apparente a la lignee bihamatus-spinaculeus. On pose comme hypothese que Creagrophorus est un genre endemique a VAmerique Centrale et qu'il est apparu a la suite de la migration vers le nord de son ancetre de VAmerique du Sud, bu le groupe apparente (Scotocryptini) s'est differencie morphologiquement et ecologiquement. La lignee hamatus a ete isolee au nord de 1'isthme de Tehauntepec tandis que la lignee jamaicensis a ete isoleee au coeur de VAmerique Centrale (et s'est disperee jusqu'en Jamaique). Finalement, C. bihamatus et C. spinaculeus sont devenus des vicariants. le premier au nord, le second au sud. en raison de Vaclivite volcanique dans la region de Chiriqui. Etant donne que les Creagrophorus ont un patron de differenciation continental, ils devraient avoir une distribution couvrant de vastes regions geographiques et leur processus de speciation devrait s'etendre sur uneperiode de temps relativement longue. On definit le groupe Aglyptinus et les Creagrophorus, et on 448 Wheeler decrit les especes de Creagrophorus. On presente aussi une clef d'identification pour chacun des genres faisant partie du groupe Aglyptinus et pour les especes de Creagrophorus, ainsi que des illustraions de I'aspect general des larves et des adultes, des illustrations des principaux caracteres et des cartes montrant la distribution geographique. On designe des lectotypes pour les especes decrites par Matthews, et enfin on discute des rapports ecologiques, cladistiques et biogeographiques. TABLE OF CONTENTS Introduction 448 Materials and Methods 449 Functional Morphology . 451 Aglyptinus Association of Genera 452 Key to Genera 453 Genus Creagrophorus Matthew 453 Key to Adults of Species 456 The Species 456 Larvae 465 Ecological Relationships 468 Cladistics 469 Character Polarity 469 Cladistic Relationships 471 Zoogeography 471 Conclusions 476 Acknowledgements 476 References 476 Index 479 INTRODUCTION Small, convex, shiny beetles (Fig. 1,2) which comprise the leiodid genus Creagrophorus Matthews are distributed in Middle America, from central Mexico south to Panama, and in Jamaica in the Antilles (Peck, 1972). Their minute size and obscure ecological habits have made these beetles exceedingly rare in museum collections. During recent field investigations on tropical, lowland, terrestrial carabids for Terry L. Erwin (National Museum of Natural History) in Panama, I collected a large series of adults and larvae of an undescribed species and, perhaps more significantly, information about the ecological relationships of Creagrophorus. Taxonomy of Creagrophorus has remained largely neglected since the generic description by Matthews in 1888. Peck (1977a) pointed out inconsistencies between the tarsal formula of his Jamaican species and the formula recorded by Matthews for his species. Following Peck's suggestion, I have re-examined Matthews' types and found that they, as well as Peck's species, have 3-3-3 tarsi. Creagrophorus, together with Aglyptinus and the Scotocryptini apparently form a monophyletic group which I define below as the taxonomically informal Aglyptinus association of genera. Members of this lineage were used as the out-group for determination of character polarity in Creagrophorus. Lack of specimens has made decisions about cladistic and zoogeographic relationships difficult. However, sister group relationships are hypothesized using existing information about structure and ecology of Creagrophorus and related genera, and a few cursory observations are made about Creagrophorus zoogeography, based on the cladistic conclusions. Revision and cladistics of Creagrophorus 449 Figures 1 - 2. Creagrophorus spinaculeus, habitus: Fig. 1, dorsal aspect; Fig. 2, lateral aspect. The purposes of this study are to define the genus Creagrophorus, revise known species, discuss ecological, cladistic (intrageneric and suprageneric), and zoogeographic relationships, and describe the larvae of a member of Creagrophorus. MATERIALS AND METHODS Materials This study is based on about 65 adult and larval Creagrophorus, including type-specimens for three previously described species. Specimens of the following species were studied as out-group taxa, and voucher specimens placed in the National Museum of Natural History (USNM, listed below): Scotocryptus sp., Aglyptinus laevis (LeConte), Aglyptinus matthewsi Champion, Aglyptinus sp. (Panama), and a new species of Aglyptinus to be described elsewhere. The following acronyms represent collections from which material was borrowed, or into which type material is deposited: BMNH British Museum (Natural History), London; CNCI Canadian National Collection of Insects, Ottawa; MCZC Musem of Comparative Zoology, Cambridge; OSUC Ohio State University, Columbus; QDWC Author's private collection, Columbus; USNM National Museum of Natural History, Washington. Quaest.Ent. 1979,15(4) 450 Wheeler Methods I have generally followed those methods discussed in a previous leiodid study (Wheeler, 1979), including both philosophical and procedural methods. Exceptions, modifications, and additions to those methods are discussed below. Species criteria I use both the theoretical and practical criteria for defining and recognizing species here as in my study of Anisotoma
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